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Meyan) Cinsinin Revizyonu

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Meyan) Cinsinin Revizyonu T.C. SELÇUK ÜNİVERSİTESİ FEN BİLİMLERİ ENSTİTÜSÜ TÜRKİYE’DE YAYILIŞ GÖSTEREN GLYCYRRHIZA L. (MEYAN) CİNSİNİN REVİZYONU Özlem ÇETİN DOKTORA TEZİ Biyoloji Anabilim Dalı Nisan-2015 KONYA Her Hakkı Saklıdır ÖZET DOKTORA TEZİ TÜRKİYE’DE YAYILIŞ GÖSTEREN GLYCYRRHIZA L. (MEYAN) CİNSİNİN REVİZYONU Özlem ÇETİN Selçuk Üniversitesi Fen Bilimleri Enstitüsü Biyoloji Anabilim Dalı Danışman: Prof. Dr. Ahmet DURAN 2015, 305 Sayfa Jüri Prof. Dr. Ahmet DURAN Prof. Dr. Hüseyin DURAL Prof. Dr. Yavuz BAĞCI Doç. Dr. Esra MARTİN Doç. Dr. Ekrem DÜNDAR Bu çalışmada, ülkemizde doğal olarak yayılış gösteren Glycyrrhiza L. (Fabaceae) cinsi taksonları morfolojik, sitotaksonomik, palinolojik, moleküler ve nümerik çalışmalara dayalı veriler kullanılarak revizyonu gerçekleştirildi. Tüm taksonların daha kullanışlı teşhis anahtarları ve kapsamlı betimleri yapıldı. Ayrıca habitat özellikleri, yayılış haritaları ve fotoğrafları verildi. Hayat formu ve IUCN kategorileri değerlendirildi. Glycyrrhiza taksonlarının kromozom sayıları belirlendi. G. asymetrica türünün somatik kromozom sayısı 2n=14, Glycyrrhiza glabra, G. echinata subsp. echinata, G. echinata subsp. macedonica, G. flavescens subsp. flavescens, G. flavescens subsp. antalyensis, G. asymmetrica taksonlarının somatik kromozom sayısı ise 2n=16’dır. Görüntü Analiz Sistemi (Bs200ProP) aracılığı ile karyotip analizleri yapıldı. Tohum yüzey ornemantasyonu taramalı elektron mikroskobu ile incelendi. Ayrıca polenler ışık ve elektron mikroskobu ile incelendi. Farklı lokalitelerden toplanan örneklerinden DNA izolasyonları yapıldı. ISSR verileri, ITS ve trnL-F bölgelerine ait sekans dizileri kullanılarak filogenetik analizleri yapıldı ve dendrogramları oluşturuldu. Glycyrrhiza cinsinin nümerik sınıflandırılmasında ise morfolojik çalışmalardan sağlanan veriler NTSYS programında analiz edilerek fenogramları oluşturuldu. Morfolojik, mikromorfolojik ve moleküler çalışmalardan sağlanan veriler Glycyrrhiza cinsinin üç alt cins (subgen. Glycyrrhiza, subgen. Liquiritia, subgen. nov. Glycyrrhizopsis) olarak ayrılmasını destekledi. Glycyrrhiza echinata subsp. macedonica taksonu Türkiye Florası için yeni kayıt olarak eklendi. Glycyrrhiza iconica hibrit bir bitki olarak tespit edildi. Beta amyrin-11 oxidase geninin intron ve ekzon bölgelerini belirlemek için Glycyrrhiza glabra ve Glycyrrhiza echinata subsp. echinata türleri kullanıldı. RNA izolasyonları yapıldıktan sonra cDNA sentezlendi. cDNA ve genomik DNA örnekleri Beta amyrin oxidase geni için sentezlenmiş primerler ile çoğaltıldı ve sekans analizi yapıldı. Yine Glycyrrhiza taksonlarında türler arasındaki polimorfizmi belirlemek amacıyla da intron ve ekzon karışımı bir bölge seçilerek PCR çalışmaları iv yapılmıştır. Bu bölge değerlendirilerek filogenetik bir ağaç oluşturulmuştur. Beta amyrin oxidase gen dizisi ile oluşturulan filogenetik ağaç ISSR, ITS ve trnL-F verileri ile oluşturulan ağaçlar ile korelasyon göstermektedir. Anahtar Kelimeler: Glycyrrhiza, ITS, ISSR, kromozom, Fabaceae, Leguminosae, polen, Sem, β-amyrin 11-oxidase, revizyon v ABSTRACT Ph.D THESIS REVISION OF THE GENUS GLYCYRRHIZA L. DISTRIBUTED IN TURKEY ÖZLEM ÇETİN THE GRADUATE SCHOOL OF NATURAL AND APPLIED SCIENCE OF SELÇUK UNIVERSITY THE DEGREE OF DOCTOR OF BIOLOGY Advisor: Prof. Dr. Ahmet DURAN 2015, 305 Pages Jury Prof. Dr. Ahmet DURAN Prof. Dr. Hüseyin DURAL Prof. Dr. Yavuz BAĞCI Assoc. Prof. Dr. Esra MARTİN Assoc. Prof. Dr. Ekrem DÜNDAR In this study, a revision of Glycyrrhiza L. (Fabaceae) taxa naturally distributed in Turkey was conducted with respect to morphology, sitotaxonomy, pollenology, molecular biology and numerical taxonomy. More practical diagnosis keys and comprehensive descriptions were also accomplished along with habitat properties, distribution maps and photographs. Life forms and IUCN categories were reviewed and revised. The number of chromosomes of Glycyrrhiza L. species was determined as G. asymetrica had 2n = 14 while the , Glycyrrhiza glabra, G. echinata subsp. echinata, G. echinata subsp. macedonica, G. flavescens subsp. flavescens, G. flavescens subsp. antalyensis, G. asymmetrica displayed 2n = 16. Karyotype and idiogram analyses were conducted through Image Analysis System (Bs200Pro). Seed surface ornamentations and pollens were investigated via electron microscope while the pollens were also studied under light microscope. ISSR results and ITS and trnL-F sequences of the taxa were used to construct phylogenetic trees. For numerical taxonomy, morphological features were scored and converted to phenograms using NTSYS program. The data obtained from morphology, micromorphology and molecular analyses suggested Glycyrrhiza organize in three sub genera as subgen. Glycyrrhiza, subgen. Liquiritia, subgen. nov. Glycyrrhizopsis. Glycyrrhiza echinata subsp. macedonica was added new record for flora of Turkey. Glycyrrhiza iconica was determined as a hybrid plant. Glycyrrhiza glabra and Glycyrrhiza echinata were used to determine the intron size of Beta-amyrin-11 oxydase. For this, cDNA synthesis following total RNA isolation were conducted. Beta amyrin-11 oxydase fragments from genomic DNA and cDNA samples were amplified via nested primers desinged for this purpose. The vi amplified fragments were sequenced using the same primers used for amplification. To determine the polymorphism of beta amynrin oxydase among Glycrrhiza species, a region of the gene including introns and exons was amplified from all the species and sequenced. The sequences were utilized to construct a phylogenetic tree which were in accordance with the trees generated using ISSR and ITS data. Keywords: chromosome, Glycyrrhiza, ITS, ISSR, Fabaceae, Leguminosae, pollen, SEM, β-amyrin 11-oxidase vii ÖNSÖZ Doktora tez çalışmam esnasında her zaman yardımlarını ve desteklerini esirgemeyen değerli danışman hocam Prof. Dr. Ahmet DURAN’a teşekkür ederim. Arazi çalışmalarında ve örneklerin temini konusunda yardım eden başta danışman hocam Prof. Dr. Ahmet DURAN’a, Doç. Dr. Ramazan Süleyman GÖKTÜRK’e, Doç. Dr. Bekir DOĞAN’a, Yard. Doç. Dr. Özge ÇETİN’e, Uzman Mustafa ÇELİK’e, Yasemin GÜRBÜZ’e, Naile ÇETİN’e ve Selçuk ÇETİN’e teşekkür ederim. Balıkesir Üniversitesi, Fen Fakültesi Araştırma Laboratuarında çalışmama imkan sağlayan, görüş ve deneyimlerini benden esirgemeyen Doç. Dr. Ekrem DÜNDAR’a teşekkür ederim. Ayrıca Balıkesir Üniversitesi, Fen Fakültesi Araştırma Laboratuarında yardımcı olan Araş. Gör. Dr. Görkem SÖNMEZ’e, Araş. Gör. Taner ÖZCAN’a, Özge TOK’a teşekkür ederim. Görüş ve deneyimlerinden faydalandığım Prof. Dr. Mustafa KÜÇÜKÖDÜK’e, Prof. Dr. Münevver PINAR’a ve Doç. Dr. Esra MARTİN’e, Doç. Dr. Tuncay DİRMENCİ’ye teşekkür ederim. Oxytropis örneklerinin temin edilmesinde bütün içtenliği ile yardım eden Yard. Doç. Dr. Seher Karaman ÖZKUL’a teşekkür ederim. Herbaryum materyallerinin incelenmesinde imkan sağlayan KNYA, GAZI, HUB, ANK, HUB, EGE, ISTF, ISTE, ISTO, AKDU, W herbaryumları yetkililerine teşekkür ederim. Selçuk Üniversitesi, Fen Fakültesi Bitki Biyolojisi Araştırma Laboratuvarındaki çalışmalarım esnasında benden yardımlarını esirgemeyen Doç. Dr. Evren YILDIZTUGAY’a, Elif GEZER’e, Fahim ALTINORDU’ya, Büşra TOSUN’a, Şerife ATİKER’e teşekkür ederim. Eğitim hayatım boyunca benden maddi-manevi desteğini esirgemeyen canım anneme ve babama teşekkür ederim Ayrıca,Tez çalışmalarım için maddi destek sağlayan TUBİTAK’a ve bu tez çalışmasına maddi destek sağlayan Selçuk Üniversitesi Bilimsel Araştırmalar Projeleri Koordinatörlüğü (BAP)’ne maddi desteklerinden dolayı teşekkür ederim. Özlem ÇETİN KONYA-2015 viii İÇİNDEKİLER ÖZET .............................................................................................................................. iv ABSTRACT .................................................................................................................... vi ÖNSÖZ ......................................................................................................................... viii İÇİNDEKİLER .............................................................................................................. ix ŞEKİLLER LİSTESİ ................................................................................................... xii ÇİZELGELER LİSTESİ ............................................................................................. xv SİMGELER VE KISALTMALAR ............................................................................ xvi 1. GİRİŞ ........................................................................................................................... 1 1.1. Glycyrrhiza cinsinin ülkemizdeki ve dünyadaki genel taksonomik problemleri .. 8 2. KAYNAK ARAŞTIRMASI ..................................................................................... 10 2.1. Fabaceae familyasının dünyadaki ve ülkemizdeki durumu ................................. 10 2.2. Papilionoideae alt familyası ve Galegeae tribusu hakkında genel bilgi .............. 22 2.3. Glycyrrhiza cinsi hakkında genel bilgi ................................................................ 25 2.3.1. Glycyrrhiza cinsinin dünyadaki ve ülkemizdeki genel durumu ................... 25 2.3.2. Glycrrhiza cinsinin biyolojik aktivitesi ile ilgili yapılan çalışmalar ............. 29 2.3.3. Glycyrrhiza cinsi üzerine yapılan moleküler çalışmalar ............................... 34 2.3.4. Fabaceae familyası ve Glycyrrhiza cinsi üzerine yapılan mikromorfolojik çalışmalar ................................................................................................................ 36 2.3.5. Glycyrrhiza cinsi üzerine yapılan karyolojik
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